34 RESTORATION POTENTIAL IN PALUSTRE VOL. 118

Source Locality Effects on Restoration Potential in Sphagnum palustre L. from 3 Ohio Sites

TONY MILLER1, Department of Biological Sciences, Kent State University, Kent, OH, USA and RANDALL J. MITCHELL, Department of Biology, The University of Akron, Akron, OH, USA.

ABSTRACT. Understanding whether propagules from different donor sources differ in their performance at a site may be important for restoration of many habitats. This study aimed at evaluating source effects inSphagnum palustre L, a peatland species, for potential use in a restoration setting. Tamarack Bog, a remnant peatland in Bath Township, Ohio, is being restored. One goal is to increase Sphagnum coverage. This study focused on the dominant species of peat moss at the bog, S. palustre. To test for source effects,S. palustre and water samples were collected from 3 different locations (Mentor Marsh, Tamarack Bog, and Singer Lake) and used in 2 experiments. performance was assessed by measuring growth in length and increase in mass. In the first experiment, a full factorial test was conducted: moss sampled from each location was grown directly in water collected from each location. In the second experiment—also a full factorial test—moss sampled from each location was separately grown on a uniform, commercially harvested, peat substrate and supplied with water collected from each location. In the design of both experiments, local adaptation would be indicated by better performance (both experiments measured length change and mass change, plus capitulum counts in the second experiment) for grown in their home water source than for plants grown in water from other sites. Ultimately, the study team did not observe evidence for local adaptation in these experiments. However, there were strong plant source effects in both experiments and some indication of differences in response to the water from different sources. Interpreting these results from a restoration standpoint, using donor plants from several source sites may improve the success of restoration.

Publication Date: November 2018 https://doi.org/10.18061/ojs.v118i2.6354 OHIO J SCI 118(2):34-42

INTRODUCTION Native plant restorations often involve the Local adaptation may be especially important in introduction of propagules from other sites to initiate understanding the biology and restoration of peatlands. or augment local populations (Hufford and Mazer Peatland restoration relies on reestablishment of 2003). In any one particular site, propagules from appropriate vegetation, including particularly one source may be best; at a different site, another Sphagnum moss. Sources of propagules for Sphagnum source may be superior. This pattern may reflect reestablishment are typically chosen because of local adaptation of plant populations, an important similar plant species composition and environmental concept in conservation (Leimu and Fischer 2008). conditions to those of the proposed restoration site Local adaptation occurs when an organism has higher (Quinty and Rochefort 2003), and because the sites are fitness in its home habitat than in others (Williams geographically nearby (Gorham and Rochefort 2003). 1966; Kawecki and Ebert 2004). Local adaptation These decisions capitalize on the fact that Sphagnum can be tested by growing individuals from different can sometimes perform better when grown in pH populations of the same species together at a site or in conditions most similar to those of the collection a common garden; finding that individuals perform site (Såstad et al. 1999). In many plant species, local better under the conditions of their home site provides populations may have as much as 50% higher fitness evidence for local adaptation (Williams 1966; Kawecki than foreign populations (Hereford 2009). Therefore, and Ebert 2004; Leimu and Fischer 2008). However, studying source effects may also be useful in the the differences in fitness are not always associated with restoration of peatlands. genetic differences, such as the case with tolerance of Sphagnum often occurs along chemical and physical sulfur compounds in the Sphagnum of the Southern gradients (Andrus 1986), and local adaptation is one Pennines (Lee and Studholme 1992), so local proposed mechanism explaining differential success adaptation can be difficult to detect inSphagnum . along the gradients. pH is one important gradient 1Address correspondence to Tony Miller, Department of © 2018 Miller and Mitchell. This article is published Biological Sciences, 256 Cunningham Hall, Kent State under a Creative Commons Attribution 4.0 International University, Kent, OH 44242. Email: [email protected] License ( https://creativecommons.org/licenses/by/4.0/) OHIO JOURNAL OF SCIENCE T. MILLER AND R.J. MITCHELL 35 determining the species distribution of Bog to stabilize the remaining habitat, and help the in peatland habitats (Vitt and Chee 1990; Rydin degraded margins recover to their original state. One and Jeglum 2013). Different haplotypes ofS. fuscum goal of the project is to improve Sphagnum coverage. (Schimp.) Klingrr. were found to occur in different Transplanting Sphagnum from donor sites may be microtopographic and pH conditions (Gunnarsson necessary to achieve this goal. The study sought to et al. 2007). Mikulášková et al. (2014) performed better understand the potential of Sphagnum source a principal coordinates analysis of S. warnstorfii to affect transplant success. This work involves S. genotypes and, of the 20 ecological variables measured, palustre, the Sphagnum species that is most common the only variable correlated with the genetic structure in the Tamarack Bog (personal observation). of the populations was pH (ranging from 4.3 to 7.2). Three source populations were chosen for this However, numerous other ecological factors may study; all contained S. palustre, but were intended affect responses. A study of S. magellanicum found to provide differing environments forSphagnum that niche differentiation was driven by differences growth in terms of pH and light. The first location in light intensity, abundance of vascular plants, air was the Tamarack Bog in Bath Township, Ohio. This humidity, and water table depth (Yousefi et al. 2017). site is categorized as a “poor fen,” with substantial More research on local adaptation is needed to groundwater inflow and outflow and a circumneutral examine the causes and consequences of home and pH (6.5 to 7; Mezentseva 2015). The site is dominated away differences (Kawecki and Ebert 2004). Here, by wetland shrubs such as Alnus incana, Vaccinium we examine this topic for Sphagnum palustre L. This corymbosum, and Toxicodendron vernix that provide species occurs in a variety of pH conditions, ranging shaded conditions in the understory across the fen. from acidic (≈3.9) to circumneutral (7.0; Andrus 1986; Water levels are within ≈10 cm of the peat surface Hájková and Hájek 2007). Since pH gradients are throughout the year (Miletti et al. 2005). important components of variation within peatlands The second location was a red maple (Acer rubrum) (Rydin and Jeglum 2013), it would be beneficial to swamp located in Mentor, Ohio (part of the Mentor determine whether the different populations of S. Marsh complex; Bernstein 1981). At this location, S. palustre have differing localized water preferences. palustre was growing on hummocks built up around This question was posed: Does water from different the roots of dead and living A. rubrum. Also, at this collecting sites affect S. palustre? To determine if source site was an encroaching population of Phragmites effects are important forS. palustre, plants from 3 australis, which, along with the red maples, provided source populations were grown in water collected mainly shaded conditions. The pH of the water at this from each location. If there are source effects, some site has not been previously reported in the literature. S. palustre populations will perform well regardless of At Mentor Marsh, the swamp habitat had flooded growth conditions. Evidence for source effects would conditions throughout the year. encourage project managers to focus donor collections The third location was an acidic kettle bog, Singer on sites with especially vigorous plants. Evidence Lake Bog (hereafter referred to as Singer Lake), against source effects would support collection of located near Green, Ohio. This location is an open, donor material for restorations from a broader range floating mat dominated by Sphagnum and leatherleaf of environmental conditions. (Chamaedaphne calyculata) with some poison sumac (Toxicodendron vernix) providing sparse shade. The METHODS AND MATERIALS S. palustre for the current study was collected from a Site Characteristics hummock that receives sunlight throughout the day. Located in the Bath Nature Preserve in Bath The water level at Singer Lake appears to be steady Township, Ohio, a tamarack peatland (Tamarack throughout the year, so the collected Sphagnum was on Bog; lat 41°10´37˝N, long 81°38´35˝W) is shrinking a hummock between 10 to 20 cm above the water table. because drainage ditches, established in the 1960s For collections at each site, field identifications were (Miletti et al. 2005), have greatly changed the hydrology confirmed using a microscope and identification key and allowed invasive species to establish. Such peatland (Crum 1984) in the lab. All from each site were destruction is common and is a leading cause of collected from a single hummock, and the water was wetland loss in Ohio (Andreas and Knoop 1992). collected from the nearest pool (within 1 meter of the Restoration efforts are now underway at the Tamarack hummock). Rainfall in the month leading up to water 36 RESTORATION POTENTIAL IN SPHAGNUM PALUSTRE VOL. 118 collection was average or below average at each of the Akron greenhouse, as earlier trials had shown that sites, so water conditions should be representative of direct sunlight in these conditions led to desiccation. typical conditions. The design yielded 3 trays of cups that were rotated Two controlled experiments investigated whether biweekly. Greenhouse temperatures were set at the S. palustre from different sites responded differently 22 °C with a natural photoperiod. to water sources. In experiment 1, the moss from each At the end of the experiment, the length of each source was grown directly in water from different shoot and the mass of the combined shoots were sources. In experiment 2, the moss was grown on measured. When the experiment ended, some cups a peat substrate and watered with water from those only contained 4 shoots. For this reason, and for each same sources. response variable, the mean value per shoot per cup was used in the analysis. Two of the collected water sources Experiment 1 occasionally harbored duckweed (Lemna minor), which To test for local adaptation, S. palustre and source was removed. Also at the end of the experiment, some water were collected from each of the study’s locations shoots had fragmented into unmeasurable pieces, between June 24 and July 15, 2015. The water sources resulting in final measurements using the mean of 4 from Singer Lake and Tamarack Bog were collected on shoots. June 24 and 25, respectively. Samples from Mentor Marsh were collected on July 15. An amount of 3.79 Experiment 2 liters (1 US gallon) of water was collected from each A second experiment was conducted to assess the location and then stored in the refrigerator (dark, effect of each water source on each Sphagnum source. 4 °C). Twenty-four hours after collection, pH was The design of the experiment entailed testing a full measured with a temperature-compensated pH meter factorial of moss and water collected from the same (VWR® SympHony® B10P). Nitrate, sulfate, and 3 locations as in experiment 1, plus another water chloride concentration were also evaluated using ion source (reverse osmosis water) as a control. As the chromatography (Dionex™ DX-100; Dionex Corp mosses collected from the field were found growing (now Thermo Scientific™); Sunnyvale, California). The at or above the water surface, a standardized substrate Sphagnum-collected at the same time as the water-was of commercially-harvested peat (Mosser Lee Co., stored in sealed clear-plastic bags in the refrigerator Millston, Wisconsin) was used to provide a substrate until the beginning of the experiment; storage times for the mosses to grow on (different than experiment 1, were between 1 to 3 weeks. The trials ran for 30 days where the mosses were grown directly in water). Water beginning in late July 2015. and moss were collected in October 2015 using the The first method of testing source effects was same sources and methods as for experiment 1. A total modeled after Ingerpuu and Vellak (2013). Beginning of 37.9 liters (10 US gallons) of water was collected in July 2015, S. palustre from each location was grown from each of the 3 locations and subsequently stored in clear-plastic cups (radius = 5.5 cm, height = 6.5 in refrigerated, opaque, tubs at the experiment location cm) for 30 days in a greenhouse (ranging between at The University of Akron greenhouse. Water pH and 22 to 24 °C). The experiment was designed as a full nutrient (nitrate, sulfate, and chloride) levels were factorial of the 3 water source treatments and the 3 measured after an initial 24-hour refrigeration period. S. palustre source locations-with 9 replicates-for a In preparation, the commercial peat was first soaked total n = 81 cups. For each cup, 5 shoots of S. palustre in reverse osmosis water for 3 days and then placed from a particular location (with capitulum intact) into 120 individual porous-plastic pots (8 cm high were each cut to 1.5 cm length, and their combined × 6 cm long × 6 cm wide). S. palustre from each site fresh mass was measured following Ingerpuu and was cut into uniform 0.5-cm-long fragments (without Vellak (2013; i.e., pressing shoots between paper capitula) and held in separate piles. Fragments were cut towels for 3 seconds before weighing). The cups from the top 10 cm of the Sphagnum shoots, necessary were initially watered with 15 mL of water from because fragments from farther down on the shoot the appropriate source, and the fill line was marked produce lower regeneration potentials in some species on each cup. The cup was filled to the mark every (Campeau and Rochefort 1996). Monday, Wednesday, and Friday. The cups were Eight fragments of S. palustre, all from the same placed on trays in deep shade in The University of source, were weighed (providing an initial fresh mass OHIO JOURNAL OF SCIENCE T. MILLER AND R.J. MITCHELL 37 measurement) and placed into one of the previously a weekly basis. Additionally, any herbaceous plants prepared peat-filled pots. This was repeated for emerging in the pots were carefully removed. n = 120 pots: 40 containing exclusively Singer Lake Experiment 2 was completed after 90 days. The moss, 40 containing exclusively Mentor Marsh moss, growth of S. palustre in each of the 120 pots was and 40 containing exclusively Tamarack Bog moss. determined by measuring final shoot length, mass Groups of 6 pots-2 each with moss from each of the change, and number of capitula. To obtain length 3 sources-were then placed together into 1 open- measurements, a plastic ruler was placed into the peat topped, clear plastic shoebox (34.6 × 21.0 × 12.4 and the longest shoot was measured from the peat cm); resulting in a total of 20 identically configured surface to the head of the capitulum. The final fresh shoeboxes. Next, 4 shoeboxes-each one labeled to mass for each pot was measured, and the percent receive water from 1 of the 4 different sources- change in mass was then obtained by dividing the were then placed together into a 110-liter clear change in mass by the initial fresh mass. plastic container; resulting in a total of 5 identically configured clear plastic containers. Ultimately, the Analysis final configuration of experiment 2 (all 120 total pots) JMP® Pro 12 was used to run statistical tests was enclosed within these 5 identically configured, on the above experiments. For water chemistry, an 110-liter clear plastic containers: each container analysis of variance (ANOVA) was used to evaluate housing 2 replicates of the full factorial of water/ differences among sites and water collection dates. In moss combinations. Later, during analysis, data on each of the plant growth experiments, full factorial the 2 replicates in each of the 110-liter containers ANOVAs were used to observe effects caused by water were averaged, yielding n = 5 for each water/moss location, Sphagnum location, and the interaction of combination. these 2 categories. For post-hoc contrasts, we used The experiment commenced in October 2015 and the Tukey’s test. was planned for 90 days. During the experiment, the 6 pots in each shoebox were watered twice a RESULTS week using 1 of the 4 different water treatments: Water Results the same source water being consistently applied to Water chemistry varied greatly among sites and each shoebox at each watering. During watering each collection dates (Table 1). The pH differed among shoebox was filled to the level of the peat surface, collecting sites (p < 0.01), being highest in the allowing the common water source to flow into and Tamarack Bog and lowest in Singer Lake on both saturate the peat in each pot. dates. Nitrate differed by collecting site and collecting Environmental conditions were monitored and date (p < 0.0001 for each). The nitrate was lowest in controlled. Existing research (Bugnon et al. 1997; Singer Lake on both dates and, in July, highest in Price 1997; Price et al. 1998; Rochefort et al. 2003) Mentor Marsh. However, in October, all 3 sites had and trials in the greenhouse indicate that maintenance similar and very low nitrate levels. Sulfate differed of a moist microenvironment and high humidity only by collection site (p < 0.001) with Mentor is important in the establishment of Sphagnum. Marsh having the highest sulfate levels. The chloride Therefore, the lids of the 110-liter containers were differed by collecting site and date (p < 0.01 for placed to maintain the humidity. To guard against each) with Mentor Marsh having higher chloride any microhabitat variations in the greenhouse, the levels. Chloride levels were higher at each site in the 110-liter containers were systematically rotated on October collection compared to the July collection. Table 1 Water chemistry for the 3 source locations Site pH Nitrate (mg/L) Sulfate (mg/L) Chloride (mg/L) Jul 2015 a Oct 2015 b Jul 2015 a Oct 2015 b Jul 2015 a Oct 2015 b Jul 2015 a Oct 2015 b Tamarack Bog 7.24 6.95 7.44 4.15 15.37 17.29 2.13 4.61 Mentor Marsh 6.27 6.80 19.84 3.72 97.02 81.65 78.00 120.53 Singer Lake 5.70 5.85 0 0.62 6.72 2.88 3.55 14.19 a n = 1; experiment 1. b mean of n = 2; experiment 2. 38 RESTORATION POTENTIAL IN SPHAGNUM PALUSTRE VOL. 118

Table 2 ANOVA values for experiment 1 (error DF a = 72) Source Number of F ratio p-value F ratio p-value DF a (length) (length) b (mass change) (mass change) b Sphagnum source 2 10.6853 <0.0001 10.5358 <0.0001 Water source 2 43.2483 <0.0001 15.8299 <0.0001 Sphagnum × water source 4 4.4797 0.003 1.2475 0.3 a Degrees of freedom. b Bold type in column indicates values of p < 0.05.

Experiment 1 the advantage to growing in water from Singer Lake Length change in experiment 1 differed nearly 10- is significant, and that growth in water from Mentor fold across all moss/water combinations (Fig. 1A), Marsh and Tamarack Bog was not distinguishable. over 2-fold among Sphagnum sources, and nearly For the moss source effect, shoots originating from 3-fold among water sources (p < 0.0001; Table 2). The the Tamarack Bog lost significantly more mass than interaction of water and Sphagnum source was also those collected from Mentor Marsh and Singer Lake significant (p = 0.003). Every combination of Sphagnum (Tukey’s comparison). Mentor Marsh and Singer Lake source and water source gained some amount of length, mosses were indistinguishable in terms of percentage but shoots from all sources performed best in Singer gain (or loss) in mass. Lake water (Fig. 1A). Moss from the Tamarack Bog performed notably worse than the others in both Experiment 2 Singer Lake and Tamarack Bog waters-but all 3 In experiment 2, the only significant cause of mosses performed similarly in Mentor Marsh water. variation for any response variable was Sphagnum Responses to water source were similar for moss from source, with water source and interaction not significant Singer Lake and Mentor Marsh, tending to be highest in all cases (Table 3). Percent gain in mass-averaged in Singer Lake water, and similarly low in the other across all water sources-was nearly 20% higher for sources. While the increase in length of the Tamarack Tamarack Bog moss than Singer Lake moss (Figure Bog moss was not significantly different in the Singer 2B; Table 3). The number of capitula produced was Lake and Mentor Marsh water sources, the Tamarack higher in Sphagnum originating from Mentor Marsh Bog moss actually performed the worst (of any moss/ and Tamarack Bog than in the Sphagnum originating water combination) in water from its home source from Singer Lake (Fig. 2C); Mentor Marsh moss and (Tamarack Bog). Tamarack Bog moss having, on average, ≈1.5 more Percent change in mass varied significantly with capitula than Singer Lake moss (Table 3; Sphagnum Sphagnum source and water source. Moss only gained source; p = 0.0002). Final length was not significantly mass in water from Singer Lake, and shoots from the affected by either Sphagnum source or water source Tamarack Bog lost mass in all water sources (Fig. 1B). (Figure 2A; Table 3), although in 3 of 4 growth For the water source effect, Tukey’s test confirms that conditions, the Tamarack Bog moss grew longest.

Table 3 ANOVA values for experiment 2 (error DF a = 48) Source Number F ratio p-value F ratio p-value F ratio p-value of DF a (length) (length) (mass (mass (capitula (capitula change) change) b number) number) b Sphagnum 2 1.2323 0.30 3.4603 0.04 10.4616 0.0002 Water 3 0.2308 0.87 0.8766 0.46 1.5773 0.21 Sphagnum × water 6 0.3778 0.89 0.1028 0.99 0.7287 0.63 a Degrees of freedom. b Bold type in column indicates values of p < 0.05. OHIO JOURNAL OF SCIENCE T. MILLER AND R.J. MITCHELL 39

FIGURE 1. S. palustre (A) length change and (B) percent change in mass in experiment 1 (error bars represent SE; n/bar = 9 cups)

DISCUSSION (5.70). However, S. palustre is commonly found in areas Water Chemistry and Experiment 1 ranging in pH from 4.0 to 7.0 (Rydin and Jeglum 2013; Singer Lake water was more conducive to S. palustre although see also Andrus (1986) which suggests pH growth than the Tamarack Bog or Mentor Marsh > 6.5 is marginal for S. palustre). With the pH of the water sources. The Singer Lake water source was the Tamarack Bog water (pH = 7.24) being only slightly only source that produced an increase in length and above this range-and with a similar change in mass an increase in mass of Sphagnum in experiment 1. in the Mentor Marsh (pH = 6.27) and Tamarack Bog The other 2 water sources resulted in decreased mass. waters-pH is less likely to be a potential driver of the Referring to Table 1, there are a few possibilities that observed differences. could explain this difference. One possible explanation Another possible explanation is the concentrations for the better performance in the Singer Lake water of the different nutrients. Increases in nitrate, sulfur source involves pH. Clymo (1973) found that some compounds, and chlorides have all been shown to Sphgnum species (S. palustre was not tested) cannot reduce Sphagnum growth (Ferguson et al. 1978; Press withstand experimental increases in pH. The Singer et al. 1986; Wilcox 1986). Singer Lake water had Lake water source was the lowest pH of the 3 wetlands less nitrate and sulfate than the other 2 sites. As for 40 RESTORATION POTENTIAL IN SPHAGNUM PALUSTRE VOL. 118

FIGURE 2. S. palustre (A) final length, (B) percent change in mass, and (C) number of capitula produced in experiment 2 (error bars represent SE; n/bar = 5 replicates) chloride, it was highest in the Mentor Marsh water found S. palustre in sites ranging from 20.04 to 90.18 (July = 78.00 mg/L). This coincides with the history mg/L (Wassen et al. 1988). The data presented here of Mentor Marsh. This site is fed by one stream that cannot distinguish among these potential causes, but has experienced increases in chloride content because it is a useful focus for future research. of nearby mining (Bernstein 1981). Yet another When comparing change in mass-and across all possible explanation is calcium. Increases in calcium water sources-the moss collected from Mentor Marsh can negatively impact Sphagnum growth (Clymo and Singer Lake outperformed the moss collected 1973). In November 2015, water samples were taken from the Tamarack Bog in experiment 1. In fact, the for measurements of calcium. The calcium level was Tamarack Bog moss of experiment 1 ended up losing, lowest at Singer Lake (4.01 mg/L) and highest at on average across all water sources, 20% of its mass. Mentor Marsh (64.13 mg/L); Tamarack Bog water This is a troublesome finding that suggests a lack of (32.06 mg/L) was intermediate between Singer Lake vigor for the S. palustre population at the Tamarack and Mentor Marsh. Calcium was in a different range Bog: Even when grown in the water from Singer Lake than found in a study from the Netherlands, which and Mentor Marsh, this moss still lost mass. Also, OHIO JOURNAL OF SCIENCE T. MILLER AND R.J. MITCHELL 41 the shoots (from all sites) grown in the Tamarack case. Furthermore, in contrast to the observations in Bog and Mentor Marsh water sources appeared more experiment 1, there was no observed shoot loss or frail than the shoots grown in the Singer Lake water color change across the water source treatments in source. In 7 of the 81 cups (and occurring in moss experiment 2. One explanation for the differing results from all sites and in water from the Tamarack Bog between the 2 experiments could be the filtering ability and Mentor Marsh) the fragile shoots broke apart of peat. One area where peat has been shown to be an into unrecognizable pieces, and this led to the loss of effective filter is in wastewater settings (Farnham and a shoot. Additionally, shoots in at least 10 of the 81 Brown 1972; Lens et al. 1994). cups (and occurring in moss from all sites) turned a grey-black color when grown in the Tamarack Bog or Implications for Restoration Mentor Marsh water sources. These 2 factors (shoot loss The results of the greenhouse experiments provide and color change) may indicate stressful conditions. information for bogs and fens undergoing restoration. Consistent with this, the Tamarack Bog and Mentor First, the S. palustre in this experiment showed no Marsh water sources had the highest measured pH evidence of local adaptation. This suggests that and calcium levels, respectively. it is reasonable to make collections from a wide array of donor sites, as opposed to a narrow choice Experiment 2 among source sites with similar vegetation and water In experiment 2, the source of S. palustre was the conditions. This could be tested with a field transplant only significant influence on plant growth; a result in study. Secondly, the S. palustre collected from Mentor contrast to experiment 1. TheS. palustre from Mentor Marsh and Tamarack Bog formed significantly more Marsh and Tamarack Bog grew more capitula, and capitula than the moss collected from Singer Lake. produced a higher percent increase of mass, than This provides evidence for the potential of vegetative S. palustre from Singer Lake. This is promising for reproduction (Cronberg 1992) because new shoots successful restoration, as capitulum development has arise when a capitulum forms from a branch coming been useful for measuring the regeneration success of off the main stem. Sphagnum. (Campeau and Rochefort 1996). As for the moss at the Tamarack Bog restoration Interestingly, water source had no significant effect site, it lost mass in each water treatment in experiment on moss growth in the peat substrate (experiment 2). 1. This could simply reflect limits of growing directly When moss was grown directly in the water of the in water with no substrate, or a problem with growth Tamarack Bog in experiment 1, the moss from all sites dormancy at that time of year. It could also indicate lost mass on average and some shoots from each source that the moss is not as healthy; however, given the were killed. However, when grown on a substrate and results of experiment 2, this does not seem likely. watered with the Tamarack Bog source (experiment The findings of this study identify 2 key points 2), the mosses grew at similar rates as with the other for restoration. First, with respect to growth, moss water sources. One reason for this could be the seasonal collections from any site would work for restoring differences in growth for this species. The growing the Tamarack Bog; however, restoration could be season for S. palustre goes from spring to July, with better fostered by using mosses with higher capitulum the moss dying back from August on (Sobotka 1974). production. Second, when spreading diaspores at Moss for experiment 1 was collected in June and July, the restoration site, they should be spread on a peat when mosses may have been at the end of their growing substrate that is less susceptible to flooding and season, so they may not have grown well in the cups. standing water. Experiment 2 started in October, when mosses may have been emerging from late-summer dormancy and ACKNOWLEDGEMENTS were therefore able to grow with more vigor. Supported in part by Crowland Ltd., the Ohio Another interesting contrast between the 2 Biological Survey Small Grants Program, and the experiments was the behavior of the moss collected Department of Biology of The University of Akron. from the Tamarack Bog. In experiment 1, the S. Thanks to the Cleveland Museum of Natural History palustre of the Tamarack Bog had-in 5 of 6 cases-the and Bath Township Parks for permission to collect least growth (or greatest loss) in terms of mass gain from the different sites, and to Jim Toppin and Diane and length gain. When grown on a peat substrate, Lucas for help in Sphagnum identification. Jean Marie in experiment 2, the S. palustre of the Tamarack Bog Hartman (Rutgers University) also provided advice had the highest percent change in mass gain in every and commented on an earlier draft of the manuscript. 42 RESTORATION POTENTIAL IN SPHAGNUM PALUSTRE VOL. 118

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